Cyrtodactylus hamidyi, Riyanto & Fauzi & Sidik & Irham & Kurniawan & Ota & Okamoto & Hikida & Grismer, 2021

Riyanto, Awal, Fauzi, Muhammad Alif, Sidik, Irvan, Irham, Mohammad, Kurniawan, Nia, Ota, Hidetoshi, Okamoto, Taku, Hikida, Tsutomu & Grismer, L. Lee, 2021, Another New Bent-toed Gecko, genus Cyrtodactylus Gray 1837 (Squamata: Gekkonidae), from Borneo, Zootaxa 5026 (2), pp. 286-300 : 289-297

Cyrtodactylus hamidyi sp. nov.

zoobank.org:act: 5CA3E52B-1D86-4D22-9F93-ACEDDFA69BEE

Recommended English common name: Hamidy’s Bent-toed Gecko

Recommended Indonesia common name: Cecak Jarilengkung Hamidy

( Figs. 2 , 3 , 4 , 5 , 6 )

Holotype. Adult male, MZB.Lace.8501 ( T062 ) ( Fig. 2 ), collected from Samaenre Semaja village , Sei Menggaris district, Nunukan regency, East Kalimantan, Indonesia (N 04.27767; E 117.22990) by I. Sidik and Mulyadi on 29 May 2011.

Paratypes (n = 6). MZB.Lace.8498, 8499 and 8500, same data as holotype ( Fig. 2 ). BORNEENSIS 09192, 09221, and 22149 ( Fig. 3 ), collected from Tawau Hills National Park , Tawau, Sabah, Malaysia by M. Matsui, K. Nishikawa, and T. Shimada on 31 July to 5 August 2004 .

Diagnosis: Cyrtodactylus hamidyi sp. nov. can be distinguished from all other congeners by the combination of the following characters: maximum SVL of at least 63 mm; no tubercles on dorsal surface of upper arm; tubercles present on ventrolateral body folds; 28–30 paravertebral tubercles; 17–20 longitudinal dorsal tubercle rows; 39–46 ventral scale rows; 17–19 subdigital lamellae on fourth toe; precloacal pit present in males; 5–7 precloacal pores in male arranged in wide Λ-shaped, absent in females; no enlarged transverse median subcaudal; enlarged median row of transverse scales; and a pair of dark semilunar-shaped markings on the upper nape.

Description of the holotype. An adult male, 55.7 mm SVL, 63.8 mm TL; head moderate in length (HL/ SVL 0.29), wide (HW/HL 0.61), slightly flattened (HH/HL 0.40), distinct from neck, triangular in dorsal profile ( Fig. 4A ); loreal scales slightly concave posteriorly, flat anteriorly; frontal and prefrontal regions concave; canthus rostralis rounded; snout elongate (SN/HL 0.40); eye large (OD/HL 0.29, OD/EyeEar 1.05); ear opening oval, almost half size of orbit diameter (EyeEar/OD 0.40), obliquely oriented ( Fig. 4B ); rostral scale rectangular, incompletely divided dorsally by a Y-shaped shallow groove; two enlarged supranasals separated from each other by three intersupranasals, two of them in contact with the smaller snout scales and one of them completely surrounded by the larger scales ( Fig. 4C ); naris oval, bordered by rostral anteriorly, anterodorsally by supranasals, and ventrally by first supralabials; 12/12 (R/L) rectangular supralabials extending to the upturn of the labial margin, tapering abruptly directly below midpoint of eye; first supralabial largest; 10/11 (R/L) infralabials extending to the upturn of the labial margin, tapering abruptly directly below midpoint of eye; rostral and loreal scales approximately same size as scales on top of head, occiput, and canthus rostralis; no tubercles on occiput or interorbital region; 36/34 (R/L) supracilliary scales elongate, smooth, largest anteriorly.

Mental triangular, bordered laterally by first infralabials and posteriorly by left and right first postmentals which contact medially for approximately 20% of their length; second postmentals seperated from each other by seven granular scales ( Fig. 4D ); small, flat gular scales gradually grading posteriorly into larger, flat, smooth pectoral and ventral scales.

Body moderate in length (AGL/SVL 0.44); ventrolateral body folds with tubercles ( Fig. 5A ); dorsum with small ganular scales interspersed with semi regularly arranged low tubercles; tubercles extend from nape to anterior of tail, exactly on the base in line the posterior of hemipenal pockets; approximately 17 longitudinal tubercle rows; 29 paravertebral tubercles. Ventral scales distinctly larger than dorsal scales, smooth, round, subimbricate, largest posteriorly; 39 ventral scale rows.

Forelimbs, dorsal surface of upper arms covered by granular scales with size approximally twice to three times larger than granular scales on dorsum; no tubercles on dorsal surface of upper and forearms. Hind limbs, dorsal surface of thigh and tibial covered by small garnular scales; tubercles present both on dorsal surface of thigh and tibia. Enlarged precloacal scales present; precloacal pit present with 6 pores arranged wide Λ-shaped ( Fig. 5B ); enlarged post-precloacal scales present ( Fig. 5B ); enlarged femoral scales absent; plantar surfaces and hind limbs granular, juxtaposed; number of lamellae under fingers (R/L): I (15/15), II (18/17), III (17/17), IV (16/15), V (11/13); number of lamellae under toes: I (13/12), II (15/15), III (17/17), IV (19/19), V (17/17).

Tail regenerated, tail length 63.9 mm; dorsal surface of original tail covered by small granular scales arranged in neat rows across dorsal tail, tubercles only at the base of the tail in line with the posterior of hemipenal pockets ( Fig. 6A, B ); ventral surface of original tail covered by smooth cycloid scales, imbricate, without transverse median subcaudal scales, median subcaudal seen larger that scales on edge ( Fig. 6C, D ).

Colouration in preservative. Ground color of head, trunk, and tail brown; a pair of semilunar dark-brown markings on posterior of occiput; two large dark-brown crossbands between axilla and groin, bordered by white network that form a vertebral line. The anterior dark band is U-shaped, extending to the eyes; ground color of ventral surface of head, trunk, hindlimbs and forelimbs whitish.

On the dorsal part of original tail from base to border of regenerated part, there are six dark bands which alternate with white bands ( Fig. 6A ). On the ventral between the base to the borderfof regenerated part of the tail there are eight dark crossbands, alternating with seven white crossbands ( Fig. 6C ).

Variation. The paratypes resemble the holotype in coloration and there seems to be no sexual dimorphism in coloration. Between axila and groin, all paratypes have a single large dark brown band forming a U-shape that extends to the eyes and is followed posteriorly by two large dark-brown bands. Two or three white networks are present between the dark bands but unlike the holotype, it does not form a vertebral line. Dorsal surface of original tail with 7 to 11 white bands. Males have precloacal pores; females do not. Detailed variation of mensural and meristic characters are presented in Table 1.

Natural history. All specimens were collected when active at the night. The holotype and paratypes from Nunukan (MZB.Lace.8498, 8499 and 8500) were collected 1 m from a stream perched approximately 30 cm above the substrate on dead wood in a logged portion of the forest, at around 1900 hrs at local time. Overall, the forest is still in good condition. The paratypes from Tawau were collected under similar conditions. BORNEENSIS 09192 was collected near riverside, on a log approximately 30 cm above the ground. BORNEENSIS 09221 was collected along a trail near headquarters, on the tree 1 m above the ground. BORNEENSIS 22149 was collected on a leaf, 50 cm above the ground. Following Grismer et al (2021), those habitat characteristics best match the “general” habitat category.

Etymology. The specific epithet hamidyi is in reference to the Indonesian herpetologist, Dr. Amir Hamidy. He has spent time a lot of teaching herpetology to the younger generations in Indonesia as well as making significant contributions to the conservation of the Indonesian herpetofauna and its diversity. He also is one of the founders of the Indonesian herpetological scociety and today is its president.

Comparison. Among the recognized species occurring in Borneo, Cyrtodactylus hamidyi sp. nov. is morphologically most like C. matsui Hikida, 1990 , in having tubercles along ventrolateral folds, lacking enlarged femoral scales, having a precloacal pit, lacking white reticulated pattern on occiput, lacking enlarged transverse median subcaudal, overlapping in number of paravertebral tubercles (28–30 versus 29), and overlapping in number of precloacal pores (5–7 versus 7–8). It differs from C. matsui in its smaller maximum SVL (65.8 versus 105 mm), in lacking tubercles on upper arm as opposed to having tubercles on upper arms,and more ventral scale rows(39–46versus 48–51). Diagnostic character comparisons to the new species from its Bornean congeners are presented in Table 2.

It is differentiated from C. baluensis in its smaller maximum SVL (65.8 versus 95 mm), in lacking tubercles on occiput as opposed to having tubercles on occiput, in lacking enlarged femoral scales as opposed to having enlarged femoral scales, in lacking enlarged transverse median subcaudals as opposed to having enlarged transverse median subcaudals, and fewer paravertebral tubercles (28–30 versus 47–60), and fewer precloacal pores (5–7 versus 9–10). It differs from C. cavernicolus Inger & King, 1961 , in its smaller maximum SVL (65.8 versus 85 mm), in having a precloacal pit as opposed to a groove, fewer ventral scale rows (39–46 versus 51–58), and fewer subdigital lamellae on the fourth toe (17–19 versus 22–26). It differs from C. consobrinus ( Peters, 1871) , in its much smaller maximum SVL (65.8 versus 125 mm), in lacking a white reticulated pattern on occiput as opposed to having white reticulated pattern on occiput, in absence of tubercles on dorsal surface of upper arm as opposed to presence, in lacking of enlarged femoral scales as opposed to having enlarged femoral scales, in lacking of enlarged transverse median subcaudals as opposed to having enlarged transverse median subcaudals, fewer ventral scale rows (39–46 versus 58–71), fewer precloacal pores (5–7 versus 9–10), and fewer subdigital lamellae on the fourth toe (17–19 versus 22–28). It differs from C. hantu Davis, Das, Leache, Karin, Brennan, Jackman, Nashriq, Chan & Bauer, 2021 , in lacking tubercles on occiput as opposed to having tubercles on occiput, having precloacal pit as opposed to a groove, and fewer paravertebral tubercles (28–30 versus 37–48). It differs from C. ingeri Hikida, 1990 , in lacking tubercles on occiput as opposed to having tubercles on occiput, in lacking enlarged transverse median subcaudals as opposed to having enlarged transverse median subcaudals, and fewer subdigital lamellae on the fourth toe (17–19 versus 23– 29). It differs from C. limajalur Davis, Bauer, Jackman, Nashriq & Das, 2019 , in its smaller maximum SVL (65.8 versus 94 mm), in having tubercles on ventrolateral body folds as opposed to lacking, in lacking of enlarged femoral scales as opposed to present, in lacking of enlarged transverse median subcaudals as opposed to having enlarged femoral scales, and more dorsal tubercle rows (17–18 versus 11–13). It differs from C. malayanus in its smaller maximum SVL (65.8 versus 93 mm), in absence of tubercles on the upper arms as opposed to presence, in lacking tubercles on occiput as opposed to having tubercles on occiput, in lacking enlarged transverse median subcaudal as opposed to present, and fewer ventral scale rows (39–44 versus 58–62). It differs from C. miriensis in lacking tubercles on occiput as opposed to having tubercles on occiput, having a precloacal pit as opposed to a groove, and fewer paravertebral tubercles (29–30 versus 47–57). It differs from C. muluensis Davis, Bauer, Jackman, Nashriq & Das, 2019 , in its smaller maximum SVL (65.8 versus 88 mm), in lacking tubercles on occiput as having tubercles on occiput, in having tubercles on ventrolateral body folds as opposed to lacking, in lacking of enlarged femoral scales as opposed to having, in having a precloacal pit as opposed to groove, in lacking of enlarged transverse median subcaudals as opposed to having enlarged transverse median subcaudals, and more dorsal tubercle rows (17–20 versus 13–15). It differs from C. pubisulcus Inger, 1958 , in having a precloacal pit as opposed to groove. It differs from C. yoshii Hikida, 1990 , in its smaller maximum SVL (63 versus 96 mm), in lacking tubercles on occiput as opposed to having tubercles on occiput, in absence of tubercles on dorsal surface of upper arm as opposed to presence, fewer ventral scale rows (39–46 versus 50–58), and fewer subdigital lamellae on the fourth toe (17–19 versus 25–30).

Furthermore, we also compare the new species to the other congeners occurs in eastern Indonesia region especially from the Wallacean region. Tubercles on upper surface of arms are absent in Cyrtodactylus hamidyi sp. nov., whereas in C. batik Iskandar, Rachmansah & Umilaela, 2011 , C. darmandvillei ( Weber, 1890) , C. halmahericus ( Mertens, 1929) , C. nuaulu Oliver, Edgar, Mumpuni, Iskandar & Lilley, 2009 , C. spinosus Linkem, McGuire, Hayden, Setiadi, Bickford & Brown, 2008 , C. tahuna, Riyanto, Arida & Koch, 2018 , C. tanahjampea Riyanto, Hamidy & McGuire, 2018 , and C. wallacei Hayden, Brown, Gillespie, Setiadi, Linkem, Iskandar, Umilaela, Bickford, Riyanto, Mumpuni & McGuire, 2008 , present. The subcaudal scales are not transversely enlarged in C. hamidyi , a condition that is shared with C. celatus Kathriner, Bauer, O’Shea, Sanchez & Kaiser, 2014 , C. fumosus ( Müller, 1895) , C. gordongekkoi ( Das, 1994) , C. jellesmae ( Boulenger, 1897) , C. halmahericus , C. laevigatus Darevsky, 1964 , C. nuaulu , C. novaeguinea ( Schlegel, 1837) , C. spinosus , C. tahuna , C. tambora Riyanto, Mulyadi, McGuire, Kusrini, Febylasmia, basyir & Kaiser, 2017 , C. tanahjampea , and C. wetariensis ( Dunn, 1927) . In contrast, transversely enlarged subcaudals are present in C. batik , C. boreoclivus Oliver, Krey, Mumpuni & Richards, 2011 , C. darmandvillei , C. hitchi Riyanto, Kurniati & Engilis, 2016, C. rex Oliver, Richards, Mumpuni & Rösler, 2016 , and C. wallacei . By lacking of spinose tubercles along ventrolateral body folds, the new species easily distinguished from C. nuaulu and C. spinosus .

Finally, we compare Cyrtodactylus hamidyi sp. nov. to congeners occurring in western Indonesia, especially from the Greater Sundas. The subcaudal scales are not transversely enlarged in C. hamidyi , a condition that is shared with C. jatnai Amarasinghe, Riyanto, Mumpuni & Grismer, 2020 , C. lateralis (Werner, 1896) , C. marmoratus Gray, 1831 , C. petani Riyanto, Grismer & Wood, 2015b , C. psarop Harvey, O’Connell, Barraza, Riyanto, Kurniawan & Smith, 2015 , C. rosichonarieforum Riyanto, Grismer & Wood, 2015a , C. semiadii Riyanto, Bauer & Yudha, 2014 , C. semicinctus Harvey, O’Connell, Barraza, Riyanto, Kurniawan & Smith, 2015 , and C. quadrivirgatus Taylor, 1962 . In contrast, transversely enlarged subcaudals are present in C. hikidai . Enlarged femoral scales are absent in C. hamidyi sp. nov., whereas in C. agamensis ( Bleeker, 1860) , C. jatnai , C. marmoratus , C. petani , C. psarops , C. semicinctus , C. quadrivirgatus , present. It differs from C. rosichonarieforum in having enlarged precloacal scales. By lacking of spinose tubercles along ventrolateral body folds, the new species is easily distinguished from C. lateralis .

Figures

FIGURE 2. Type series of Cyrtodactylus hamidyi sp. nov. from Nunukan, East Kalimantan, Indonesia. Bars = 5 mm.

FIGURE 3. Paratypes of Cyrtodactylus hamidyi sp. nov. from Tawau, Sabah, Malaysia. Bars = 5 mm.

FIGURE 4. Head features of holotype (MZB.Lace.8501) of Cyrtodactylus hamidyi sp. nov. (A) Dorsal view. (B) Lateral view. (C) Dorsoanterior view showing rostal region, white arrows indicate inter nares. (D) Ventral view showing mental region, white arrows indicate small scales between two second post-mentals. Scale not shown.

FIGURE 5. Lateral trunk and precloacal region features of holotype (MZB.Lace.8501) of Cyrtodactylus hamidyi sp. nov. (A) Tubercles present along long ventrolateral body folds, indicated by red arrows. (B) Six precloacal pores arranged in wide Λ-shaped (indicated by red arrows) and presence of enlarged post precloacal scales (under black circle), and femoral region without distinct enlarged femoral scales. Scale bars = 5 mm.

FIGURE 6. Tail features of holotype (MZB.Lace.8501) of Cyrtodactylus hamidyi sp. nov. (A) Dorsal surface of original tail covered by small granular scales arranged in neat rows across dorsal portion of tail, no tubercles except at the base in line with the posterior of hemipenal pockets. (B) Insert of dorsal surface. (C) Ventral surface of original tail covered by smooth cycloid scales, imbricate, without transverse median subcaudal scales, median subcaudal seen larger that scales on edge. (D) Insert of ventral surface. Scale bar = 5 mm.